A Revision of the Scaly Toadfishes, Genus Batrachoides, with Descriptions of Two New Species from the Eastern Pacific

Batrachoides differs from all other American genera of Batrachoididae in having the body covered with scales and in having two subopercular spines. Nine species are recognized: liberiensis (Steindachner) from the Gulf of Guinea (with beninensis Regan a synonym); manglae Cervigón from Venezuela; gilberti Meek and Hildebrand from the Caribbean side of the Yucatán Peninsula south to Panamá; surinamensis (Bloch and Schneider) from Honduras to Salvador, Brazil; goldmani Evermann and Goldsborough from Atlantic-draining streams in the Río Usumacinta and Rio Grijalva in México and Guatemala; boulengeri Gilbert and Starks from the Bay of Panamá; pacifici (Günther) from Panamá to northern Peru; waltersi n.sp. from Acapulco, Mexico south to the Gulf of Nicoya, Costa Rica; and walkeri n.sp. known only from the holotype from the Bay of Panamá. B. boulengeri and B. surinamensis are a pair of Pacific-Atlantic geminate species as are B. pacifici and B. gilberti. B. waltersi is clearly related to B. boulengeri and B. surinamensis but has fewer dorsal and anal fin rays and vertebrae. B. walkeri is similar to B. boulengeri and B. surinamensis in small eye size and high vertebral and dorsal ray counts but differs from both in having reduced squamation on the head. Attempts to analyze relationships within the genus led to an unresolved trichotomy between B. liberiensis, the B. boulengeri species group (including B. surinamensis, B. waltersi, and B. walkeri), and the B. pacifici group (including B. gilberti, B. manglae, and B. goldmani). Brief notes are included on swimbladder morphology, fecundity, food habits, and parasites.